early Homo

Holy crap 2015 was a big year for fossils. And how fortuitous that 2016 begins on a Fossil Friday – let’s recap some of last year’s major discoveries.

Homo naledi

Some Homo naledi mandibles in order from least to most worn teeth.

The Homo naledi sample is a paleoanthropologist’s dream – a new member of the genus Homo with a unique combination of traits, countless remains belonging to at leasta dozen individuals from infant to old adult, representation of pretty much the entire skeleton, and a remarkable geological context indicative of intentional disposal of the dead (but certainly not homicide, grumble grumble grumble…). The end of 2015 saw the announcement and uproar (often quite sexist) over this amazing sample. You can expect to see more, positive things about this amazing animal in 2016.

We’ll be presenting a bunch about Homo naledi at this year’s AAPA meeting in Hotlanta. I for one will be discussing dental development at Dinaledi- here’s a teaser:

As long as we’re talking about the AAPA meetings, my colleague David Pappano and I are organizing a workshop, “Using the R Programming Language for Biological Anthropology.” Details to come!

Lemur graveyard

Homo naledi wasn’t the only miraculously copious primate sample announced in 2015. Early last year scientists also reported the discovery of an “Enormous underwater fossil graveyard,” containing fairly complete remains of probably hundreds of extinct lemurs and other animals. As with Homo naledi, such a large sample will reveal lots of critical information about the biology of these extinct species.

Australopithecus deyiremeda

We also got a new species of australopithecus last year. Australopithecus deyiremeda had fat mandibles, a relatively short face (possibly…), and smaller teeth than in contemporaneous A. afarensis. One tantalizing thing about this discovery is that we may finally be able to put a face to the mysterious foot from Burtele, since these fossils come from nearby sites of about the same geological age. Also intriguing is the possible evidence, based on published CT images (above), that A. deyiremeda had relatively advanced canine and delayed molar development, a pattern generally attributed to Homo and not other australopithecines (if this turns out to be the case, you heard it here first!).

Lomekwian stone tool industry

Roughly contemporaneous with A. deyiremeda, Harmand et al. (2015) report the earliest known stone tools from the 3.3 million year old site of Lomekwi 3 in Kenya. These tools are a bit cruder and much older than the erstwhile oldest tools, the Oldowan from 2.6 million years ago. These Lomekwian tools, and possible evidence for animal butchery at the 3.4 million year old Dikika site in Ethiopia (McPherron et al. 2010; Thompson et al. 2015), point to an earlier origin of lithic technology. Fossils attributed to Kenyanthropus platyops are also found at other sites at Lomekwi. With hints at hominin diversity but no direct associations between fossils and tools at this time, a lingering question is who exactly was making and using the first stone tools.

Earliest Homo

The reconstructed Ledi Geraru mandible (top left), compared with Homo naledi (top right), A. deyiremeda (bottom left), and the Uraha early Homo mandible from Malawi (bottom right). Jaws are scaled to roughly the same length from the front to back teeth; the Uraha mandible does not have an erupted third molar whereas the others do and are fully adult.

Just as Sonia Harmand and colleagues pushed back the origins of technology, Brian Villmoare et al. pushed back the origins of the genus Homo, with a 2.7 million year old mandible from Ledi Geraru in Ethiopia. This fossil is only a few hundred thousand years younger than Australopithecus afarensis fossils from the nearby site of Hadar. But the overall anatomy of the Ledi Geraru jaw is quite distinct from A. afarensis, and is much more similar to later Homo fossils (see image above). Hopefully 2016 will reveal other parts of the skeleton of whatever species this jaw belongs to, which will be critical in helping explain how and why our ancestors diverged from the australopithecines. (note that we don’t yet have a date for Homo naledi – maybe these will turn out to be older?)

The earlier hominin fossil record wasn’t the only part to be shaken up. A small molar (KNM-ER 51261) and a set of associated hip bones (KNM-ER 5881) extended the lower range of size variation in Middle and Early (respectively) Pleistocene Homo. It remains to be seen whether this is due to intraspecific variation, for example sex differences, or taxonomic diversity; my money would be on the former.

At the later end of the fossil human spectrum, researchers also announced an archaic looking mandible dredged up from the Taiwan Straits, and a more modern-looking brain case from Israel. The Penghu 1 mandible is likely under 200,000 years old, and suggests a late survival of archaic-looking humans in East Asia. Maybe this is a fossil Denisovan, who knows? What other human fossils are waiting to be discovered from murky depths?

The Manot 1 calvaria looks very similar to Upper Paleolithic European remains, but is about 20,000 years older. At the ESHE meetings, Israel Hershkovitz actually said the brain case compares well with the Shanidar Neandertals. So wait, is it modern or archaic? As is usually the case, with more fossils come more questions.

Crazy dinosaurs

Yi qi was bringing Skeksi back, and its upper limb had a wing-like shape not seen in any other dinosaur, bird or pterosaur. There were a number of other interesting non-human fossil announcements in 2015 (see here and here), proving yet again that evolution is far more creative than your favorite monster movie makers.

What a year – new species, new tool industries, new ranges of variation! 2015 was a great year to be a paleoanthropologist, and I’ll bet 2016 has just as much excitement in store.

We’re learning about the divergence between robust Australopithecus and early Homo 2.5-ish million years ago in my Human Evolution class this week. Because of this multiplicity of contemporaneous species, when scientists find new hominin fossils in Early Pleistocene sites, a preliminary question is, “What species is it?”

To help my students learn how we know whether certain fossils belong to the same species, and to which group new fossils might belong, in this week’s lab we compared tooth sizes of Australopithecus boisei and early Homo. After seeing how tooth sizes differed between these groups, students then tested whether they could determine whether two “mystery” fossils (KNM-ER 60000 and 62000; Leakey et al. 2012) belonged either group.

Early Pleistocene hominin fossils from Kenya. Left to right: KNM-ER 406, ER 62000 and ER 1470. At the center is one f the lab’s “mystery jaws.”

The first purpose of this lab was to help familiarize students with skull and tooth anatomy of early Pleistocene humans. Although lectures and readings are full of images, a lab activity forces students to spend time visually examining fossils. Plus, they’re in 3D which is a whole D greater than 2D – the visual equivalent of going to eleven! The second goal of the lab was to help prepare students for their term projects, in which they must pose a research question about human evolution, generate predictions, and find and use data to test hypotheses.

If you’re interested in using or adapting this activity for your class, here are the handout and data sheet into which students enter their measurements. The data sheet specifies the fossils that can be downloaded from africanfossils.org. Some relevant fossils (i.e., KNM WT 15000 and ER 992) were not included because the 3D scans yield larger measurements than in reality.

Figure 3 from Ward et al. 2015. A little distal to the hip, yes, but the pun still works. Views are, going clockwise starting at the top the top left, from above, from below, from the back, from the side, and from the front.

There’s also a partial ilium associated with the femur – that makes a pretty complete hip!

Figure 5 from Ward et al. shows the fossil. Jump for joy that it’s complete enough for us to tell it comes from the left side!

Despite how fragmentary the femur and ilium are, the researchers were able to estimate the diameter of the femur head and hip socket reliably. The hip joints are smaller than all Early Pleistocene Homo except for the Gona pelvis. Comparing ER 5881 the large contemporaneous KNM-ER 3228 hip bone, the authors found these two specimens to be more different in size than is usually seen between sexes of many primate species. The size difference best matches male-female differences in highly dimorphic species like gorillas.

Ward et al. find that the specimen generally looks like early Homo but that the inferred shape of the pelvic inlet is a little different from all other Early and Middle Pleistocene human fossils. The authors take this discrepancy to suggest that there was more than one “morphotype” (‘kind of shape’), and therefore possibly species, of Homo around 1.9 million years ago. While I wouldn’t just yet go so far as to say this anatomy is due to species differences, I do agree that KNM ER 5881 helps our understanding and appreciation of anatomical variation in our early ancestors. Like all great fossil discoveries, the more we find, the more we learn that we don’t know. Here’s to more Homo hips in the near future!

Last year I brought up the implications of the small female pelvis from Gona, Ethiopia for body size variation in Homo erectus (see previous post). This individual was much smaller than other Middle Pleistocene Homo fossils, indicating size variation comparable to highly sexually dimorphic gorillas and unlike recent human populations. Before this pelvis, most known Homo erectus fossils were fairly large (comparable to living people), with only a few hints of much smaller individuals (e.g., KNM-ER 427000, KNM-OL 45500). Now joining this petite party, this tiny troop, this little lot, this compact cadre, etc., is KNM-WT 51261, a 750,000 year old molar from Kenya (Maddux et al., in press).

Occlusal area for first molars in the genus Homo. The tooth image is from Fig. 2 and the plot from Fig. 3 in Maddux et al. Lookit how tiny it is!

This ‘new’ specimen substantially increases the range of size variation among early African H. erectus molars, although the expanded range isn’t remarkable compared with later Homo samples such as from Zhoukoudian cave in China or Neandertals. What is different, though, is that most of the highly variable samples show a fairly continuous range of variation, while the WT 51261 molar is a considerable outlier from the rest of the African Middle Pleistocene sample (a lot like the situation with the Gona pelvis). So this tooth re-raises an important question: were smaller specimens like Gona and WT 51261 as rare in life as they are in the fossil record, or was such great size variation common in the Middle Pleistocene? How we reconstruct what kind of animal Homo erectus was differs depending on the answer to this question.

I’m reading up on life history in Homo erectus for a few projects I’m working on, and something’s just caught my eye. A 2012 issue of Current Anthropology presents a series of papers from the 2011 symposium, “Human Biology and the Origins of Homo.” This issue is full of great stuff, and to top it all off, it can be accessed online for free! (here’s the JSTOR link)

Gary Schwartz has a paper here recounting what is known (or as he stresses, what is still largely unknown) about growth and life history in early Homo. Dental evidence accumulated over the past 30 years has pointed to a rapid (ape-like) life cycle for fossil hominins, in comparison with a slow, long and drawn out human pattern. But much of the evidence against a human-like pattern is somewhat indirect. For instance, Holly Smith (1991) has shown that there’s a pretty tight relationship between brain size and age at first molar (M1) eruption in Primates:

Fig. 1 from Schwartz (2012). “Bivariate plot of ln M1 emergence age in months (y) versus ln cranial capacity in cubic centimeters (x) for a sample of anthropoids.” The hominins and humans are the open shapes, to which I’ve visually fitted the red line.

It’s a very high correlation (r=0.98). This means that armed with simply an animal’s cranial capacity, which is fairly easy to estimate given complete enough fossils, one can estimate with a bit of confidence its likely age range for M1 emergence. With brain sizes between apes’ and ours, fossil hominins can be estimated to have erupted their M1s at younger ages than us. Many subsequent studies of tooth formation, based on the microscopic remnants of tooth development, have supported these inferences. So presumably, faster, ape-like dental development could be extrapolated to mean ape-like body growth rates and other aspects of life history as well.

But although this is a tight relationship, there are deviations. As Schwartz notes in the article, and others have noted before, high correlations found when examining large interspecific groups (e.g., primates as a whole) often break down when the focus is on smaller groups of more closely related species (e.g., just apes). Based on the relationship figured above, humans are expected to erupt M1 around 7 years of age, but nearly all humans erupt M1 closer to 6 years (hence the open diamond for humans is below the regression line). What hominins appear to share in common with humans is a younger age at M1 eruption than expected for primates of their brain sizes (the red line I’ve added to the figure).

Hominins’ faster dental development and eruption may be ape-like in absolute terms, but eruption ages may be human-like when their brain size is taken to account. As with many life history variables, the significance of this similarity (if anything) is difficult to ascertain.

Some big changes here at Lawnchair Anthropology. I just successfully defended my dissertation (Mandibular Growth in Australopithecus robustus, more info on that to come), and moved to Kazakhstan to begin my new job in the School of Humanities and Social Sciences at Nazarbayev University. I landed in Astana about 22 hours ago, so I should be asleep, battling (or succumbing to) jetlag, but some friends have pointed me to newly published early Homo fossils from Kenya, dating to between 1.9-1.6 million years ago (Leakey et al., 2012). See Adam Van Arsdale’s blog, the Pleistocene Scene, for great historical background and perspective on these new fossils.

Now, one of the themes of my dissertation is that there is lots of interesting information to be gleaned from fossils that we’ve known about for a long time (many of the A. robustus mandibles featured in my research have been known for decades). But dammit if some of these much more recently discovered fossils point to tantalizing variation in hominids just later than 2 million years ago (note I’m careful to say “variation” rather than “diversity”). In light if this variation, Adam discusses the similarities between one of these Kenyan fossils (KNM-ER 60000) and the large mandible from Dmanisi, which was discovered in only in the year 2000 (Gabunia et al., 2002).

Piggy-backing off Adam, I’d like to point out similarities between another of the new fossils, the KNM-ER 62000 face of a juvenile, and the recently discovered A. sediba juvenile face (Berger et al., 2010). These two fossils are at the same stage of dental development, so they’re roughly at the same stage of life. They are close in geological age, but A. sediba is from South Africa. Below are figures of A. sediba (left) and the ER 62000 face (right). The pics should be to scale, modified from the original publications. (sorry I couldn’t remove the background from the top left one)

What do you think? Pretty different, right? WRONG! Below I’ve superimposed the ER 62000 face onto A. sediba (slightly recolored and transparented for contrast). Remember that these are to scale.

In front view (left), the ER 62000 face is almost identical to A. sediba, right down to the positions of the teeth. THIS DOES NOT MEAN THAT I THINK THESE TWO FOSSILS REPRESENT THE SAME SPECIES. In side view, however, some differences do become apparent. Notably, the front of the A. sediba maxilla projects a bit further forward than ER 62000, and the nasal and orbital anatomy are also fairly different. THIS DOES NOT MEAN THAT I THINK THESE ARE DIFFERENT SPECIES. (although I would be surprised if these fossils turned out to be the same animal)

Leakey et al. liken these new Kenyan fossils to the cranium KNM-ER 1470, from the same region and at 1.9 million years old. But what’s weird to me is that ER 1470 actually looks a bit more like the juvenile A. sediba in the side view (as reconstucted; the face and braincase of ER 1470 are actually separated, leaving it unclear just how the two parts fit together). Here are all three specimens, to scale:

From left to right: ER 62000, A. sediba, ER 1470

Now, the ER 1470 comparison isn’t really fair – ER 1470 is an adult and it is much larger: the bottom of ER 1470’s eye socket is about the same height as the top of A. sediba‘s. The size difference is probably the main reason why its face below the nose sticks out as much as A. sediba‘s, even though the latter is smaller. (I should note, too, that theadult A. sediba mandible is superficially very similar in gonial and ramus anatomy to another of the recently published Kenyan specimens, ER 60000).

The point of all these comparisons is not to say whether these fossils are the same species, but rather to point out that there are actually striking similarities between fragmentary fossils, and it’s not clear what exactly these similarities (or differences, for that matter) mean. Maybe my eye was drawn to the ER 62000-A. sediba comparison not because of any evolutionary relationship, but because these fossils are in similar stages of growth and development – if it weren’t waaaaay past my bedtime I’d love to compare these fossils with other similarly-aged fossils (like D2700 from Dmanisi and KNM-WT 15000, also from Kenya).

All of these fossils (except ER 1470) were discovered in the past few years. I’ve said it before and I’ll repeat it now: this is a great time to study paleoanthropology.

Two views of an Acheulian handaxe adorn the cover of this week’s Nature (right). Always happy to see paleoanthropology stuff be classy, front-page news. The cover highlights Christopher Lepre’s and colleagues’ announcement of what may be the oldest Acheulian tools known.

To recap stone tools: The first good evidence of tool use by humans’ ancestors are the Oldowan lithics from the 2.6 million year old site of Gona in Ethiopia (Semaw et al. 2003). McPherron and others (2010) reported 2 possibly-cut-marked animal bones from the 3.4 million-year old site of Dikika; but this latter evidence is a bit too scant for us to really be sure our ancestors had adopted technology this early. Anyway, the Oldowan was a very basic tool industry, consisting largely of crude flakes taken off cobbles. It may sound lame, but even the most basic stone-tool-making requires some skills, trust me, it’s kinda hard. So stone tools appear roughly 2.5 million years ago, which is also about the time that we have fossils that might document the earliest members of our genus Homo. Sweet.

The legend goes that the next technological revolution doesn’t come until about 1 million years later – until around 1.5 million years ago, stone tools were quite basic. But after a while we start seeing these “handaxes” or “bifaces” (cuz flakes are removed from both of the core’s faces; see above) that have become kind of the hallmark of what’s termed the Acheulian industry. I’m sure there are other key indicators but what do I know, I’m not an archaeologist. Arguably, the rise of the Acheulian from its humble Oldowan beginnings is a milestone in human cognitive evolution – a more complex tool should require a more complex brain, right? Lepre and team announced today that they have some Acheulian handaxes from the Kenyan site of Kokiselei-4, dating to 1.76 million years ago. The authors draw two conclusions: 1) the Acheulian (and thereby more advanced cognition) is a few hundred thousand years older than previously thought, and 2) the co-occurrence of Acheulian and Oldowan tools at this time indicates the presence of contemporaneous human species with different cognitive capabilities.

Now what’s a bit odd here is that the presence of handaxes among otherwise Oldowan assemblages is not a new or unique thing. In her archaeological research at Olduvai Gorge in Tanzania, Mary D Leakey distinguished some assemblages as “Developed Oldowan.” Here’s a relevant blurb from a study by Y. Kimura (2002: 292-293):

“Leakey recognized two distinctive industries, Oldowan and Acheulian, from Bed I through Bed III at Olduvai. The former was characterized by the presence of various choppers and attributed to Homo habilis sensu lato, whereas the latter was traditionally defined to contain bifaces more than 40-60% of the tools, and attributed to H. erectus sensu lato.

The Oldowan was then classified into Oldowan (1.87-1.65 mya) and Developed Oldowan (1.65-0.6 mya) based on the increased light-duty tools, spheroids and bifaces in the latter. The Developed Oldowan coexisted with Acheulian” (emphasis mine)

So the co-occurrence of Oldowan (i.e. choppers) and Acheulian (some handaxes) is known from other sites, albeit not until around 1.5 million years ago. Too bad I’m not an archaeologist nor know more about lithics, because I wish I could put the new Kokiselei-4 assemblage into this context – just how is it different from “Developed Oldowan”? As John Hawks pointed out before I did, “developed Oldowan” doesn’t appear in the Lepre et al. discussion. Hrm. Then they make this statement:

“Homo erectus is traditionally thought to be the first hominin to disperse from Africa, yet the oldest known out-of-Africa fossil hominin sites lack stone tools or preserve only Oldowan-style artefacts. … Our data indicate that the earliest development of the Acheulian occurred in Africa at 1.76 [million years] ago and was contemporaneous with or perhaps pre-dated the earliest hominin dispersals into Eurasia (Lepre et al. 2011: 84).

They then go on to suggest that two contemporaneous species lived in Africa in the early Pleistocene – one of these species invented the Acheulian and stayed in Africa, while the other species was too dumb to make anything beyond Oldowan, and instead these dullards left Africa to colonize the rest of the world. This silly scenario seems to stem from an under-appreciation of what Dmanisi demonstrates (possibly since the recent dating paper by Reed Ferring and others only came out a few months ago, probably after the Lepre et al. paper was in press). The Dmanisi fossils establish that hominins more primitive than later Homo erectus (Rightmire et al. 2006) had dispersed into Eurasia by around 1.85 million years ago (if not earlier), with mere Oldowan technology (Mgeladze et al. 2010, Ferring et al. 2011). So Lepre et al.’s claim that the earliest Acheulian “was contemporaneous or perhaps pre-dated” the first out-of-Africa dispersals just isn’t true. And without that, there’s no support for the silly scenario of a smart, techno-savvy but stationary species being contemporaneous with a colonizing but less crafty-and-cunning species.

It’s really cool if the Kokiselei-4 tools truly represent the earliest record of the Acheulian. But, it should be clear by now that we can’t simply equate technology and taxonomy. So how old is the Acheulian and why does it matter? I’m fine with a 1.76 million year date, but I also don’t think it matters too much. (sorry to be so Dmanisi-centric)